© 2020. Published by The Company of Biologists Ltd | Biology Open (2020) 9, bio053546. doi:10.1242/bio.
053546
RESEARCH ARTICLE
‘Whip from the hip’: thigh angular motion, ground contact
mechanics, and running speed
Kenneth P. Clark1, *, Christopher R. Meng2 and David J. Stearne1
ABSTRACT
During high-speed running, lower limb vertical velocity at touchdown
has been cited as a critical factor needed to generate large vertical
forces. Additionally, greater leg angular velocity has also been
correlated with increased running speeds. However, the association
between these factors has not been comprehensively investigated
across faster running speeds. Therefore, this investigation aimed to
evaluate the relationship between running speed, thigh angular
motion and vertical force determinants. It was hypothesized that thigh
angular velocity would demonstrate a positive linear relationship with
both running speed and lower limb vertical velocity at touchdown. A
total of 40 subjects (20 males, 20 females) from various athletic
backgrounds volunteered and completed 40 m running trials across a
range of sub-maximal and maximal running speeds during one test
session. Linear and angular kinematic data were collected from
31–39 m. The results supported the hypotheses, as across all
subjects and trials (range of speeds: 3.1–10.0 m s−1), measures of
thigh angular velocity demonstrated a strong positive linear
correlation to speed (all R 2>0.70, P<0.0001) and lower limb vertical
velocity at touchdown (all R 2=0.75, P<0.001). These findings suggest
thigh angular velocity is strongly related to running speed and lower
limb impact kinematics associated with vertical force application.
KEY WORDS: Bipedal gait, Locomotor control, Sprinting
INTRODUCTION
Whether at sub-maximal or maximal intensity, maintaining a
constant forward running speed presents several mechanical
challenges. Among these requirements, steady-speed running
necessitates that the net three-dimensional acceleration of the
center of mass (COM) over an entire stride cycle is zero. Therefore,
the horizontal and lateral forces must integrate to zero during each
stride, and the average vertical ground reaction force over a complete
stride cycle must equal the body’s weight (Raibert, 1986). Although a
wealth of experimental research has investigated human locomotor
performance (Blickhan, 1989; Farley et al., 1993; McMahon and
Cheng, 1990; Nagahara et al., 2014; Rabita et al., 2015; Seyfarth
et al., 2003; Weyand et al., 2000), a complete description of the
mechanics that runners select to satisfy the demands of high-speed
running has not yet been completely established. Therefore, further
1
Department of Kinesiology, West Chester University of Pennsylvania, West
Chester, PA 19383, USA. 2Department of Chemical and Biomolecular Engineering,
University of Notre Dame, Notre Dame, IN 46556, USA.
*Author for correspondence (kclark@wcupa.edu)
K.P.C., 0000-0002-6934-1271
This is an Open Access article distributed under the terms of the Creative Commons Attribution
License (https://creativecommons.org/licenses/by/4.0), which permits unrestricted use,
distribution and reproduction in any medium provided that the original work is properly attributed.
Received 12 May 2020; Accepted 27 August 2020
experimental investigation is warranted, to explore several important
aspects of running mechanics.
Nearly a half-century of research has examined vertical force
application during running (Cavagna, 1975; Cavanagh and
Lafortune, 1980; Clark et al., 2017; Kuitunen et al., 2002; Munro
et al., 1987; Nilsson and Thorstensson, 1989; Weyand et al., 2000,
2010). As speed increases and ground contact time decreases, the
average vertical force applied during ground contact must increase
to achieve the vertical impulse (force×time) necessary to support the
body and rebound the COM into the next step. Recent research has
demonstrated that faster speeds are typified by larger vertical forces
primarily during the first half of ground contact (Bezodis et al.,
2008; Clark and Weyand, 2014; Kuitunen et al., 2002), as vertical
force application during the second half of ground contact is similar
between sprinters and non-sprinters (Clark and Weyand, 2014). The
large forces in the first half of ground contact can be explained by a
Newtonian force–motion relationship (Bobbert et al., 1991), linking
the sharp rising edge of the vertical force waveform observed at
faster speeds to the rapid impact deceleration of the lower limb upon
touchdown (Clark et al., 2017; Nigg et al., 1987).
In addition to lower-limb impact mechanics, proximal kinematic
inspection (thigh segments) may provide additional insight.
Previous publications on bipedal locomotion have suggested that
limbs function as harmonic oscillators with torsion springs at the hip
and scissor-like thigh angular motion (McGeer, 1990). In this
investigation, we present a framework of thigh angular motion
during the gait cycle (see Materials and Methods). Reciprocal
oscillations at the hip result in angular movement patterns between
the limbs, with one thigh flexing while the other extends. These
angular motions are critical for facilitating steady forward speed
(Raibert, 1986). Harmonic angular limb motion has been observed
not only in human runners (Novacheck, 1998; Sides, 2015), but also
in avian bipedal locomotion (Blum et al., 2014; Rubenson et al.,
2007) and models of running robots (McGeer, 1990; Thompson and
Raibert, 1989).
Furthermore, measurements of leg angular velocity have been
positively related to running speed in human sprinting (Belli et al.,
2002; Kivi et al., 2002; Kunz and Kaufman, 1981; Mann and
Herman, 1985; Mann and Murphy, 2018; Miyashiro et al., 2019)
and cited as a critical factor for running speed in robotic legged
locomotion (Thompson and Raibert, 1989). However, despite
prior evidence linking faster running speeds to both greater vertical
ground reaction force and leg angular velocity, the connection
between these two factors has not yet been fully explored across a
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range of speeds and runners. Theoretically, increased thigh
angular velocity is not only necessary to satisfy the kinematic
demands of faster running speeds, but may also lead to greater
lower limb vertical velocity at touchdown, which has been
established as a critical factor for generating the large mass-
specific vertical forces required for high speed running (Clark
et al., 2017; Mann and Murphy, 2018). Potentially, the angular
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RESEARCH ARTICLE Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546

velocity of the front swing thigh as it extends during the last
portion of the flight phase could be directly related to the lower
limb velocity at touchdown, serving as a contributing factor to
vertical force application and running speed.
Therefore, the aim of this study was to examine thigh angular
motion and kinematic factors influencing vertical force application
across a range of speeds. Our first hypothesis was that thigh angular
velocity would demonstrate a positive and linear relationship with
running speed across a range of sub-maximal and maximal
intensities. Our second hypothesis was that thigh angular velocity
would demonstrate a positive and linear correlation with lower limb
velocity at touchdown across a range of running speeds, lending
insight into the relationship between thigh angular motion and
vertical force determinants. These concepts may contribute to the
understanding of human running, with practical applications for
training interventions aimed at enhancing running performance.
RESULTS
Running mechanics across entire range of speeds
The subjects differed greatly in size and athletic background, but
demonstrated similar modifications to running mechanics across the
range of speeds. Representative data for thigh angular position
versus time is presented in Fig. 1, with a male recreationally trained
athlete at sub-maximal and maximal speeds displayed in Fig. 1A
and C, and a male sprinter at sub-maximal and maximal speeds
displayed in Fig. 1B and D.
Spatial–temporal variables and thigh angular kinematic variables
are presented for a single subject (male sprinter) across his individual
range of speeds in Fig. 2, and for the subject population as a whole
across all subjects and trials in Fig. 3. Spatial–temporal and thigh
angular kinematic variables across all subjects and trials (categorized
by slow, intermediate and fast speeds) are also presented in Table 1
and Table S2.
For the spatial–temporal variables, contact time (Tc) decreased
with increasing speed for individual subjects (Fig. 2A) and for all
subjects and trials [Fig. 3A; Table 1: F(2,151)=135.1, P<0.001].
Flight time (Tf ) was slightly briefer in duration at fast speeds than at
slow and intermediate speeds for all subjects and trials [Table 1:
F(2,151)=7.9, P<0.001], although the differences in Tf between
slow and intermediate speeds were not statistically significant
(Table 1: P>0.8). Step rate (SR) increased across the range of speeds
for both individual subjects (Fig. 2B) and for all subjects and trials
[Fig. 3B; Table 1: F(2,151)=179.4, P<0.0001]. Step length (SL)
increased across speeds for individual subjects (Fig. 2B) and for all
subjects and trials [Fig. 3B; Table 1: F(2,151)=67.1, P<0.0001],
although the increases in SL from intermediate to fast speeds were
not statistically significant (Table 1: P=0.17). Across all subjects
and trials, the mean distance travelled by the COM during ground
contact (Lc) was 0.97±0.01 m. However, Lc increased slightly
across speeds (Table 1: H=10.4, P<0.01), though the increases in Lc
from intermediate to fast speeds were not statistically significant
(Table 1: P>0.9). Analysis regarding vertical velocity of the lower

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Fig. 1. Thigh angular position versus time for two representative subjects, with dashed regions of the graph indicating the ground contact phase.
(A,C) Male recreationally trained athlete at sub-maximal and maximal speeds. (B,D) Male sprinter at sub-maximal and maximal speeds. Faster running
speeds were achieved with higher frequencies and greater total amplitudes of thigh angular motion, resulting in greater thigh angular velocities. At top speed,
the slope of the angular position versus time curve during ground contact was steeper for the sprinter (D) than for the recreationally trained athlete (C),
indicating a greater absolute value of the average angular velocity of the stance thigh during ground contact (ωc). Per Eqn 6, greater ωc was a direct
determinant of the faster top running speed attained by the sprinter.
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Fig. 2. Kinematic variables for a single representative subject (male sprinter) across his individual range of speeds. (A) Ground contact time, Tc.
(B) Step rate, SR, and step length, SL. (C) Total thigh excursion during ground contact phase, θc, and total thigh excursion from peak extension through peak
flexion, θtotal. (D) Average thigh angular velocity during entire gait cycle, ωavg, and lower limb vertical velocity at instant of touchdown (Ankle Vztd).
limb (ankle marker) at the instant of touchdown (Ankle Vztd) is
presented in the last section of the Results.
With respect to thigh angular kinematics, faster speeds were
generally characterized by higher frequency and greater total
amplitude of thigh angular motion (θtotal ), as illustrated by
representative data in Fig. 1. Increases in θtotal occurred with
speed for individual subjects (Fig. 2C) and for all subjects and trials
(Fig. 3C; Table 1: H=91.6, P<0.0001). Increases in θtotal across the
range of speeds were due to both increases in thigh flexion (θflex)
and increases in thigh extension (θext) (Appendix B and Table S2).
Across all subjects and trials, the total excursion angle during
contact (θc) approached one radian (mean θc=0.97±0.01 radian).
However, there were increases in θc across the range of speeds for
individual subjects (Fig. 2C) and for all subjects and trials (Fig. 3C;
Table 1: H=71.8, P<0.0001). Increases in θc across the range of
speeds were due to both increases in thigh touchdown (θtd) and in
thigh takeoff (θto) (Appendix B and Table S2).
All measures of thigh angular velocity significantly increased
across speeds. There was a positive and linear relationship between
absolute average thigh angular velocity during the entire gait cycle
(ωavg) and speed for individual subjects (Fig. 2D) and for all
subjects and trials [Fig. 3D; Table 1: F(2,151)=204.2, P<0.0001].
Additionally, the average angular velocity of the stance thigh during
ground contact (ωc) increased with speed for all subjects and trials
[ωc=1.14x−0.89, R 2=0.88, P<0.0001; Table 1: F(2,151)=257.4,
P<0.0001]. Per Eqn 4, Tc showed a strong relationship with ωc for
all subjects and trials (Tc=0.002x 2−0.045x+0.36 where x is ωc,
R 2=0.91), and Tc also had a strong relationship with ωavg for all
Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546
subjects and trials (Tc=0.62x −0.86 where x is ωavg, R 2=0.90).
Per Eqn 6, ωc× leg length (L0) showed a direct positive linear
relationship with speed for all subjects and trials with the best fit
line constrained through the origin (ωc×L0=0.94x, R 2=0.91,
P<0.0001). Also, ωretr increased with speed across all subjects
and trials (ωretr=0.80x−2.14, R2=0.71, P<0.0001; Table 1:
H=70.2, P<0.0001). Finally, across all subjects and trials, ωavg
was related to the other two measures of thigh angular velocity, as
ωavg had a strong positive relationship to both ωc (ωc=1.35x−0.55
where x is ωavg, R 2=0.92, P<0.0001) and ωretr (ωretr=0.98x−2.10
where x is ωavg, R2=0.80, P<0.0001).
Running mechanics across top speeds
Kinematic variables were analyzed across all top speed trials and
with runners grouped by top speed and sex. There were significant
differences in top speed for the fast versus slow subjects when
analyzed within sex (Table 2, males: P<0.001, Δ=14.2%; females:
P<0.01, Δ=13.2%). Faster top speeds were significantly related to
briefer Tc across all top speed trials (Fig. 4A) and when analyzing
fast versus slow runners within sex (Table 2, males: P<0.001,
Δ=21.3%; females: P<0.01, Δ=15.0%). Faster top speeds were also
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significantly related to greater SR across the entire subject
population (Fig. 4B) and when analyzing fast versus slow runners
within sex (Table 2, males: P<0.01, Δ=8.6%; females: P=0.04,
Δ=6.5%). Similar relationships were found for SL across all top
speed trials (Fig. 4B) and when analyzing fast versus slow runners
within sex (Table 2, males: P=0.002, Δ=5.7%; females: P=0.015,
Δ=6.6%). However, top speed was not significantly related to Tf
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Fig. 3. Kinematic variables for all subjects across all speeds (n=154). Best-fit equations and P-values listed here where appropriate, with R2 values
presented in accompanying panels. (A) Ground contact time (Tc=0.78x −0.89). (B) Step rate (SR=0.31x+1.58, P<0.0001) and step length (SL=0.72x 0.49).
(C) Total thigh excursion during ground contact phase (θc=−0.01x 2+0.20x+0.22) and total thigh excursion from peak extension through peak flexion
(θtotal=0.51x 0.56). (D) Average thigh angular velocity during entire gait cycle (ωavg=0.82x−0.08, P<0.0001).

(across all top speed trials: Tf=−0.03x+0.14, R 2=0.07, P=0.10;
Table 2, males: P=0.30, Δ=4.3%; females: P=0.60, Δ=2.5%) or Lc,
(across all top speeds Lc=0.01x+0.93, R 2=0.01, P=0.62; Table 2,
males: P=0.07, Δ=6.8%; females: P=0.51, Δ=2.2%).
For thigh angular kinematics, faster top speeds were
characterized by higher frequency and greater θtotal (see Fig. 1C
versus D for representative data). Across all top speed trials, θtotal
was positively and significantly related to faster top speeds
(Fig. 4C), although the differences in θtotal did not reach
significance when analyzing fast versus slow runners within sex
(Table 2, males: P=0.099, Δ=5.7%; females: P=0.057, Δ=6.2%).
For all top speed trials, θc was slightly greater than one radian
(mean θc=1.07±0.01 radian). However, θc was negatively and
significantly related with speed across all top speed trials
(Fig. 4C), and differences in θc were significant for males (but
not females) when analyzing fast versus slow runners within sex
(Table 2, males: P=0.009, Δ=6.6%; females: P=0.057, Δ=3.4%).
Additional top speed data for θtd, θto, θext and θflex are presented in
Appendix B and Table S3.
females: P=0.003, Δ=11.9%). Finally, ωretr was positively and
significantly related to faster top speeds across all top speed trials
(ωretr=1.18x−5.36, R 2=0.64, P<0.0001) and when analyzing fast
versus slow runners within sex (Table 2, males: P=0.014, Δ=28.2%;
females: P=0.011, Δ=32.5%).
Lower limb impact velocity, running speed, and thigh angular
kinematics
Ankle Vztd demonstrated positive and linear relationships with both
running speed and with measures of thigh angular velocity (ωavg and
ωretr). Ankle Vztd increased across the range of speeds for individual
subjects (Fig. 2D) and for all subjects and trials (Fig. 5A; Table 1:
H=72.1, P<0.0001). Faster top speeds were positively and
significantly related to greater Ankle Vztd across all top speed
trials (Ankle Vztd=0.29x−0.23, R 2=0.40, P<0.001), although when
analyzing fast versus slow top speeds within sex, Ankle Vztd
demonstrated significant differences for females but did not reach
significance for males (Table 2, males: P=0.118, Δ=13.9%;
females: P=0.015, Δ=19.8%). Ankle Vztd was positively and
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Across all top speed trials, ωavg was positively and significantly
related to faster top speeds (Fig. 4D) and when analyzing fast versus
slow runners within sex (Table 2, males: P=0.002, Δ=14.2%;
females: P=0.003, Δ=12.8%). Likewise, ωc was positively and
significantly related to faster top speeds across all top speed trials
(ωc=0.77x+2.57, R 2=0.58, P<0.0001) and when analyzing fast
versus slow runners within sex (Table 2, males: P=0.003, Δ=13.7%;
significantly related to ωavg across all subjects and trials (Fig. 5B),
and when analyzed for top speed trials only (Ankle
Vztd=0.37x−0.41 where x is ωavg, R 2=0.52, P<0.001). Finally,
Ankle Vztd was significantly related to ωretr across all subjects and
trials (Ankle Vztd=0.30x+0.83 where x is ωretr, R 2=0.75, P<0.0001),
and when analyzed for top speed trials only (Ankle
Vztd=0.23x+1.15 where x is ωretr, R 2=0.54, P<0.001).

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Table 1. Kinematic variables across speeds, categorized by percentage top speed

Slow <75% top speed (n=44 trials) Intermediate 75–93% top speed (n=48 trials) Fast >93% top speed (n=62 trials)
Spatial–temporal variables
Speed, (m s−1) 4.78±0.16 6.94±0.12† 8.18±0.12*†
Tc (s) 0.204±0.006 0.144±0.003† 0.122±0.002*†
Tf (s) 0.129±0.004 0.131±0.002 0.119±0.001*†
SR (steps s−1) 3.03±0.04 3.66±0.04† 4.18±0.04*†
SL (m) 1.56±0.04 1.89±0.02† 1.96±0.02†
Lc (m) 0.93±0.01 0.98±0.01† 0.99±0.01†
Ankle Vztd (m s−1) 1.30±0.06 1.81±0.05† 2.21±0.05*†
Thigh angular variables
θc (rad, deg) 0.85±0.02, 48.6±0.9 0.98±0.01†, 56.2±0.8 1.05±0.01*†, 60.3±0.5
θtotal (rad, deg) 1.20±0.03, 68.6±1.7 1.49±0.02†, 85.4±1.0 1.64±0.02*†, 94.2±0.9
ωavg (rad s−1, deg s−1) 3.80±0.12, 217.7±6.6 5.48±0.09†, 314.1±5.2 6.82±0.11*†, 390.9±6.1
ωc (rad s−1, deg s−1) 4.36±0.17, 249.8±9.8 6.92±0.12†, 396.7±6.9 8.74±0.12*†, 500.9±7.1
ωretr (rad s−1, deg s−1) 1.85±0.15, 105.9±8.9 3.29±0.17†, 188.6±9.8 4.42±0.17*†, 253.5±9.7
All angular variables are absolute values. Values are means± standard error of the mean (s.e.m.).
Spatial–temporal variables: running speed, ground contact time (Tc), flight time (Tf ), step rate (SR), step length (SL), contact length (Lc) and vertical velocity of
lower limb at instant of touchdown (Ankle Vztd).
Thigh angular variables: thigh excursion during ground contact phase (θc), thigh excursion from peak extension to peak flexion (θtotal ), average thigh angular
velocity during entire gait cycle (ωavg), average thigh angular velocity during ground contact phase (ωc), and average thigh angular retraction velocity (ωretr).
†
indicates significantly different than slow speed (P<0.05).
* indicates significantly different than intermediate speed (P<0.05).

DISCUSSION
Experimental tests of hypotheses
We undertook this investigation to explore the relationship between
thigh angular motion, ground contact mechanics and running speed.
Based on our framework, we hypothesized that thigh angular
velocity would have a strong positive linear relationship with
running speed across the entire range of trials, and across the
subjects’ range of top speeds. We also hypothesized that thigh
angular velocity would have a strong positive linear relationship
with lower limb vertical velocity at touchdown. We recruited a
heterogenous group of 40 subjects that included males and females
across a range of different sizes (height range: 1.52–1.94 m, mass
range: 45.5–117.0 kg), from a variety of athletic backgrounds
(recreationally trained individuals, intercollegiate team sport
athletes, and competitive track and field athletes), and had them
run across more than a threefold range of speeds (3.1–10.0 m s−1).
Individual subjects ran faster by increasing both the frequency
and amplitude of thigh angular motion, resulting in an increase in
thigh angular velocity (Figs 1 and 2). This same pattern was
consistent across all subjects and trials, as increases in running
speed demonstrated a strong positive relationship with thigh angular
velocity (Table 1). This was best represented by the 91% shared
variance between running speed and ωavg across all trials (Fig. 3D).
Additionally, across all top speed trials, faster top speeds had a
positive, significant relationship to both ωavg, ωretr and ωc (Table 2),
with 74% shared variance between running speed and ωavg across
top speed trials (Fig. 4D). When top speed trials were analyzed with
subjects grouped by speed and sex, all measures of thigh angular
velocity were significantly greater in fast subjects compared to their
slow counterparts. Comparing fast and slow runners within sex
(Table 2), the percentage differences in ωavg (∼13 to 14%) were
nearly identical to the differences in top speed (∼13 to 14%).
Likewise, our data also supported the second hypothesis. Across
a range of speeds from a slow jog to a maximal sprint, Ankle Vztd
demonstrated a significant positive relationship with speed (Table 1),
with 68% shared variance between running speed and Ankle Vztd

Table 2. Kinematic variables across top speed trials, with subjects categorized by sex and top speed
Fast males (n=10) Slow males (n=10) Fast females (n=10) Slow females (n=10)
Spatial–temporal variables
Top speed (m s−1) 9.50±0.10† 8.23±0.13 8.06±0.17* 7.06±0.07
Tc (s) 0.105±0.002† 0.130±0.006 0.117±0.004* 0.136±0.004
Tf (s) 0.114±0.004 0.109±0.003 0.123±0.004 0.120±0.004
SR (steps s−1) 4.58±0.06† 4.21±0.09 4.19±0.09* 3.93±0.08
SL (m) 2.07±0.03† 1.96±0.02 1.93±0.03* 1.80±0.03
Lc (m) 1.00±0.01 1.07±0.03 0.94±0.02 0.96±0.03
Ankle Vztd (m s−1) 2.47±0.15 2.15±0.10 2.16±0.12* 1.77±0.08
Thigh angular variables
θc (rad, deg) 1.02±0.02†, 58.6±0.9 1.09±0.02, 62.5±1.0 1.06±0.02, 60.7±1.0 1.10±0.02, 62.8±1.2
θtotal (rad, deg) 1.73±0.04, 98.9±2.3 1.63±0.04, 93.4±2.2 1.65±0.04, 94.3±2.2 1.55±0.03, 88.7±1.7
ωavg (rad s−1, deg s−1) 7.82±0.13†, 448.0±7.7 6.78±0.25, 388.7±14.1 6.88±0.22*, 394.4±12.4 6.05±0.10, 346.9±5.6
ωc (rad s−1, deg s−1) 9.76±0.19†, 559.1±10.8
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8.51±0.31, 487.4±18.0 9.13±0.23*, 522.8±13.0 8.10±0.20, 464.3±11.5

ωretr (rad s−1, deg s−1) 5.75±0.22†, 329.6±12.8 4.33±0.47, 248.2±26.8 4.23±0.40*, 242.5±22.9 3.05±0.11, 174.6±6.5
Spatial–temporal variables: running speed, ground contact time (Tc), flight time (Tf ), step rate (SR), step length (SL), contact length (Lc) and vertical velocity of
lower limb at instant of touchdown (Ankle Vztd).
Thigh angular variables: thigh excursion during ground contact phase (θc), thigh excursion from peak extension to peak flexion (θtotal ), average thigh angular
velocity during entire gait cycle (ωavg), average thigh angular velocity during ground contact phase (ωc) and average thigh angular retraction velocity (ωretr).
†
indicates significantly different (P<0.05), fast males versus slow males.
*indicates significantly different (P<0.05), fast females versus slow females.

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Fig. 4. Kinematic variables for all subjects across top speed trials (n=40). Best-fit equations and P-values listed here, with R2 values presented in
accompanying panels. (A) Ground contact time (Tc=−0.013x+0.23, P<0.001). (B) Step rate (SR=0.30x+1.78, P<0.0001) and step length (SL=0.10x+1.14,
P<0.0001). (C) Total thigh excursion during ground contact phase (θc=−0.03x+1.28, P=0.014) and total thigh excursion from peak extension through peak
flexion (θtotal=0.07x+1.03, P<0.001). (D) Average thigh angular velocity during entire gait cycle (ωavg=0.77x+0.58, P<0.0001).

(Fig. 5A). Similar results were evident across all top speed trials, as
Ankle Vztd was significantly related to top speed for the entire subject
population. When comparing within speed and sex, Ankle Vztd was
significantly greater for fast females than slow females (Table 2), and
although this differential did not reach significance for males, both
fast males and fast females had Ankle Vztd that were more than 13%
greater than their slow counterparts. Ankle Vztd was also significantly
related to thigh angular velocity (ωavg, and ωretr) across the range of
speeds, with 75% shared variance between Ankle Vztd and ωavg
(Fig. 5B) and between Ankle Vztd and ωretr. Similar findings were
also evident across top speed trials, as Ankle Vztd was significantly
related to both ωavg and ωretr at top speed.

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Fig. 5. Lower limb vertical velocity at instant of touchdown (Ankle Vztd) for all subjects and trials (n=154). Best-fit equations and P-values listed here,
with R2 values presented in accompanying panels. (A) Ankle Vztd versus running speed (Ankle Vztd=0.27x−0.02, P<0.0001). (B) Ankle Vztd versus average
thigh angular velocity during entire gait cycle (Ankle Vztd=0.33x+0.01 where x is ωavg, P<0.001).

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Fig. 6. Simplified planar representation of the ground contact phase. (A) Geometry of the ground contact leg, including: symmetrical θtd and θto, θc, L0
and Lc. (B) θtd and θto during the ground contact phase. This framework assumes that the leg angle (from ball of foot to hip, θtd or θto in A) is equal to the
thigh angle (θtd or θto in B, respectively).

Framework evaluation
While the data strongly supported the hypotheses, it is important to
note that alternative outcomes were possible. A strong positive
linear relationship between running speed and ωavg may not have
occurred if the fundamental framework assumptions regarding
ground contact geometry (θc and Lc) were violated. Based on prior
evidence it was assumed that θc≈1.0 radian and that Lc≈L0≈1.0 m
(Clark and Weyand, 2014; Farley et al., 1993; Gatesy and Biewener,
1991; He et al., 1991; Weyand et al., 2010). In this study, θc
increased slightly across speeds for all subjects and trials (Figs 2C,
3C, and Table 1), but the mean increase in θc from intermediate to
fast speeds was only 0.07 radians (∼4°). Likewise, Lc increased
slightly across speeds (Table 1), although increases in Lc from
intermediate to fast speeds were not statistically significant. Not
surprisingly, there was some between-subject variability in Lc, with
values differing between the shortest female and the tallest male by

Biology Open

Fig. 7. Simplified representation of thigh angular motion during the entire gait cycle. This representation assumes symmetrical anti-phase flexion and
extension values during ground contact and flight phases. Thigh angular kinematics as a function of time are determined by the parameters of frequency
(f=1/T, where T is the period) and amplitude (A). The figures in inset diagrams a–g illustrate the thigh angular position in correspondence with the angular
motion presented in the graph. The dashed regions of the graph indicate the ground contact phase, with ωc corresponding to the slope of the angular position
versus time curve from touchdown (+0.5 radian) to takeoff (−0.5 radian).

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RESEARCH ARTICLE
Fig. 8. Example graphs of thigh angular position versus time for one
limb, examining the theoretical effects of altering frequency (f=1/T) and
amplitude (A). In A–C, the gray line has f=1.33 Hz and A=0.60 radians. The
purple line illustrates the effects on thigh angular velocity that result from
altering frequency and amplitude. (A) Increasing A without altering f.
(B) Increasing f without altering A. (C) Increasing both A and f. The dashed
regions of the graph indicate the ground contact phase, with average ωc
corresponding to the slope of the angular position versus time curve. In all
three panels, the purple line has greater ωc than the gray line because of
increased A and/or f.
more than 0.2 m at top speed. However, when normalized to L0,
these values did not demonstrate extreme deviation from the
geometric assumptions, with Lc/L0 ratios of 1.03 and 1.11 at top
speed for the shortest female and tallest male, respectively. Given
that the mean θc was 0.97±0.01 radians across all subjects and trials,
and the mean Lc across all subjects and trials was 0.97±0.01 m, the
experimental data generally adhered to the assumptions presented in
the framework. Prior research has suggested that humans and other
bipedal runners are likely constrained to these excursion angles and
contact lengths due to leg extensor muscle mechanical advantage
(Biewener, 1989; Weyand et al., 2010).
Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546
A second assumption related to symmetrical thigh flexion and
extension values about the zero-axis during both ground contact and
flight phases. Our assumption of symmetrical θtd and θto angles of
0.5 radians during ground contact was not entirely supported,
although mean θtd and θto values approached 0.5 radians at
intermediate and faster speeds (Appendix B and Table S2).
Conceivably, if the θtd angles had deviated further from the
expected values, it could have weakened the correlation between
thigh angular velocity (ωavg and ωretr) and Ankle Vztd. As it related
to thigh flexion and extension during flight, recent evidence (Mann
and Murphy, 2018) has suggested that runners might exhibit more
‘front-side’ mechanics at faster speeds (i.e., more thigh flexion and
less thigh extension during the flight phase). This evidence
questions the veracity of assuming symmetrical flexion/extension
about the zero-axis during the flight phase. In our data set, θtotal
increased across the range of speeds due to increases in θflex that
were proportionately bigger than the increases in θext ( Appendix B
and Table S2). Furthermore, across the top speed trials, fast males
and fast females had θtotal that was shifted more front-side, with a
greater proportion of θtotal occurring in θflex than in θext, compared to
slow males and slow females (Fig. 1C,D, Appendix B and
Table S3).
Finally, the thigh segment motion was qualitatively oscillatory
and reciprocal. Peak flexion of one thigh generally exhibited
temporal coordination with peak extension of the other thigh,
especially at faster speeds (Fig. 1). Although the data exhibited
some deviation from exact sinusoidal motion, this was expected due
to slightly variable coordination strategies across a broad range of
speeds. This variation was not substantial enough to disrupt another
important assumption related to continuous harmonic thigh motion
throughout the gait cycle, which indicates that ωavg must increase for
ωc to increase. Our data supported this premise, as across all subjects
and trials, there was 92% shared variance between ωavg and ωc, and
90–91% shared variance between Tc and measures of thigh angular
velocity (ωavg and ωc). Therefore, despite some data not fully
conforming to simplifying assumptions, the two experimental
hypotheses were still supported strongly by the results.
Building on prior research
Several previous publications have observed the relationship
between measures of leg angular velocity and running speed (Kivi
et al., 2002; Kunz and Kaufman, 1981; Mann and Herman, 1985;
Mann and Murphy, 2018; Miyashiro et al., 2019). Furthermore,
faster running speeds have been associated with measures of hip
joint strength, power, torque, work and muscle activation (Belli
et al., 2002; Bezodis et al., 2008; Copaver et al., 2012; Deane et al.,
2005; Dorn et al., 2012; Nagahara et al., 2020; Schache et al., 2011).
Here, Figs 6–8 and Eqns 1–8 (Materials and Methods) provide a
geometric/mathematical explanation for the linear relationship
between ωavg and running speed. Data from this investigation
supports these concepts, as results indicated a direct positive linear
relationship between increases in ωavg of 1.0 rad s−1 and increases in
running speed of ∼1.2 m s−1 (Figs 3D and 4D) and Ankle Vztd of
∼0.33 m s−1 (Fig. 5B).
Contributing to the strong correlations was the measurement of
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ωavg, and not just thigh angular velocity in one specific phase of the
cycle such as flexion or extension. Measuring ωavg provided a
comprehensive representation of the oscillatory frequency and
amplitude necessary to achieve a given running speed. Although
thigh angular acceleration was not directly reported in this study, the
measurement of ωavg over the entire gait cycle accounted for the
rapid angular accelerations that occurred as the thigh reversed from
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flexion to extension or vice versa. These vigorous flexion/extension
reversals were more pronounced in faster runners than slower
runners at top speed (Fig. 1C,D), and undoubtedly contributed to
the greater ωavg observed in fast runners than their slow
counterparts.
The results presented here also provide insight into the limb
kinematics underlying force application. Numerous prior
investigations have established the relationship between high-
speed running and mass-specific vertical force (Bundle and
Weyand, 2012; Clark and Weyand, 2014; Dorn et al., 2012;
Kuitunen et al., 2002; McGowan et al., 2012; Weyand et al., 2000,
2010), with recent research indicating that greater vertical forces are
due in large part to faster vertical velocities of the lower limb at
touchdown combined with a rapid deceleration of the lower limb
during initial ground contact (Clark et al., 2014, 2017). In the
present study, the values of Ankle Vztd across a range of speeds
(Fig. 5A) closely aligned with data from recent publications (Clark
et al., 2017; Udofa et al., 2019). Additionally, although correlation
and causation must be interpreted with caution, our results suggest
that increased thigh angular velocities contribute to the greater
Ankle Vztd observed at faster speeds (Fig. 5B).
Collectively, these findings suggest that thigh angular velocity is
related to vertical force determinants. This is perhaps best illustrated in
Fig. 2D, which depicts nearly parallel increases in ωavg and Ankle Vztd
across the subject’s range of speeds. Slower speeds are characterized by
reduced thigh oscillatory frequencies and amplitudes, resulting in
relatively decreased thigh angular velocities and slower vertical
velocities of the lower limb at touchdown. Faster running speeds
require increased thigh oscillatory frequencies and amplitudes, resulting
in greater thigh angular velocities and faster vertical velocities of the
lower limb at touchdown. The latter, when combined with a stiff ground
contact and rapid lower limb deceleration upon ground contact (Clark
et al., 2017), contributes to the greater mass-specific vertical forces
required for faster running speeds (Kuitunen et al., 2002; McGowan
et al., 2012; Weyand et al., 2000).
Future considerations and practical applications
While building on prior findings, this study raises many topics
worthy of further investigation. First, we intentionally recruited a
heterogenous group of male and female subjects from a range of
sizes and athletic backgrounds, and analyzed running mechanics
across a threefold range of speeds. Undoubtedly, some of the strong
statistical relationships we found between speed and the kinematic
variables were due to this heterogenous subject pool and broad
range of speeds. However, we justified our approach based on the
desire to elucidate macro-level determinants of running speed from
slow jogging to maximal sprinting. Although further research is
necessary to confirm whether the relationships established here will
generalize to a less diverse group of athletes within more narrow
ranges of speed (i.e. elite sprinters at top speed), the existing
evidence regarding the thigh angular kinematic determinants of
speed (Mann and Murphy, 2018) suggest that our major findings
may generalize to homogenous subject populations.
Potential applications of these findings for performance
improvement in human sprinting are intriguing. Coaching cues
such as ‘whip from the hip’ have been popularized by some well-
known practitioners, emphasizing a vigorous scissor-like action of
the thighs (Bosch and Klomp, 2005), and these instructions appear
to have practical merit. Similar to a hammer striking a nail, high-
speed running requires fast rotational and tangential velocity prior to
impact combined with a stiff collision upon impact. Likewise, our
findings suggest that greater top speeds require fast thigh angular
Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546
retraction velocities in an open kinetic-chain movement prior to
ground contact, combined with a stiff stance limb that allows the
thigh to extend rapidly in a closed kinetic chain movement
throughout ground contact. We speculate that a relatively stiff
lower limb during ground contact is not only imperative for brief
contact times and large vertical forces (Arampatzis et al., 1999; Belli
et al., 2002; Clark et al., 2017; Kuitunen et al., 2002), but also to
allow the thigh to continuously extend at high angular velocities
throughout ground contact. Any excessive compliance in the lower
limb during contact may hinder rates of thigh extension, prolonging
ground contact time and decreasing running speed.
Our findings align with recent research linking faster running
speeds to increased hip joint muscular activation, torque, work and
power (Belli et al., 2002; Bezodis et al., 2008; Copaver et al., 2012;
Dorn et al., 2012; Nagahara et al., 2020; Schache et al., 2011).
However, few studies have examined methods for longitudinal
improvement of these determinants. Although the effects of lower
body wearable resistance (Macadam et al., 2017) and hip flexor
strengthening (Deane et al., 2005) have been investigated, the best
methods for enhancing thigh angular velocity during sprinting are
not clearly established, and require expanded investigation.
Conceivably, coaches and athletes could aim to enhance thigh
flexion angular velocity in open-kinetic chain movements (forward
swing phase), or thigh extension angular velocity in open-kinetic
chain (retraction) or closed-kinetic chain (ground contact)
movements. Given the coordinated and reciprocal oscillatory
motion generally demonstrated by the thigh segments, any
longitudinal improvement in thigh extension velocity should
require corresponding improvement in thigh flexion velocity, and
vice versa. In other words, increasing only thigh flexion or thigh
extension capability, without concomitant improvement in the other
variable, is likely to limit the runner because the thighs must
complete the powerful scissor-like action in synchrony. Therefore,
optimal training interventions likely need to target enhancing thigh
angular velocity in both flexion and extension actions during both
open- and closed-kinetic chain movements.
Concluding remarks
Here we investigated thigh motion and lower limb vertical velocity
at touchdown across a broad range of runners and speeds. As
hypothesized, thigh angular velocity had a direct linear relationship
to both running speed and the lower limb kinematics underlying
vertical force application. Our results suggest that increases in thigh
angular velocity are not only necessary to match the kinematic
demands for high-speed running, but also contribute to the larger
mass-specific vertical forces necessary to support faster speeds.
Therefore, interventions aimed at improving running performance
likely need to elicit an increase in thigh angular velocity through all
phases of the gait cycle.
MATERIALS AND METHODS
Framework
A simplified planar representation of the ground contact phase is depicted in
Fig. 6. This framework assumes symmetrical touchdown and takeoff angles
during ground contact and that the leg angle (from ball of foot to hip, θtd or
θto in Fig. 6A) is equal to the thigh angle (θtd or θto in Fig. 6B, respectively).
Biology Open
The Lc is determined by leg length L0 and θc:
uC
Lc ¼ 2  L0  sin : ð1Þ
2
For faster running speeds, prior research has suggested that the total
excursion angle during contact is approximately 1.0 radian for humans and
other bipedal runners (Farley et al., 1993; Gatesy and Biewener, 1991;
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He et al., 1991). Thus, for normal contact excursion angles where θc≈1.0
radian, Eqn 1 demonstrates that contact length is approximately equal to leg
length (Lc≈L0). Furthermore, since horizontal velocity during the flight
phase is constant (negating wind resistance), the runner’s forward speed is
determined by the time it takes the COM to traverse Lc, where Tc is ground
contact time (Weyand et al., 2010):
Lc
Speed ¼ : ð2Þ
Tc
Because Lc≈L0, and this distance is generally limited to about 1.0 m for
most adult human runners at faster running speeds (Clark and Weyand,
2014; Weyand et al., 2010), increases in speed are usually accompanied by
decreases in Tc (Dorn et al., 2012; Nummela et al., 2007; Weyand et al.,
2010).
Additionally, leg angular velocity is a crucial determinant of locomotor
stability and control (Seyfarth et al., 2003). Average angular velocity of the
stance thigh during ground contact (ωc) is equal to the total contact
excursion angle divided by ground contact time:
uc
vc ¼ : ð3Þ
Tc
Eqn 3 can be rearranged so that Tc is expressed as a function of θc and ωc:
uc
Tc ¼ : ð4Þ
vc
Inserting Eqn 4 into Eqn 2 and substituting Lo for Lc yields Eqn 5:
Lc L0 vc
Speed ¼ ¼ ¼ L0  : ð5Þ
Tc ðuc =vc Þ uc
Assuming a total excursion angle θc equal to 1.0 radian, running speed is
directly related to L0 and ωc:
Speed
Speed ¼ L0  vc or ¼ vc : ð6Þ
L0
Furthermore, harmonic oscillatory thigh motion has been observed
during high-speed running in humans (Novacheck, 1998; Sides, 2015),
with contralateral limbs exhibiting reciprocal anti-phase flexion and
extension during bipedal gait (Blum et al., 2014). Fig. 7 depicts a
simplified example of thigh angular motion that assumes symmetrical
anti-phase flexion and extension values during ground contact and flight
phases. Because this framework is based on equations of harmonic
motion, thigh angular kinematics as a function of time are determined by
the parameters of frequency ( f=1/T, where T is the time period)
and amplitude (A). Thigh angular position as a function of time is
displayed in Eqn 7:
uðtÞ ¼ A  sinð2p  f  tÞ: ð7Þ
Thigh angular velocity as a function of time is displayed in Eqn 8:
du
vðtÞ ¼ ¼ A  ð2p  f Þ  cosð2p  f  tÞ: ð8Þ
dt
Graphically, the ground contact phase is represented by the dashed lines
in Fig. 7, from touchdown (+0.5 radian) to takeoff (-0.5 radian). Thus, ωc
is equal to the slope of the angular position versus time curve during the
ground contact phases (slope of the dashed line in Fig. 7B,C,E and F).
The steeper the slope of the angular position versus time curve in
Fig. 7B,C,E,F, the greater the absolute value of ωc, and the faster the
running speed (normalizing for L0, per Eqn 6).
The figures and equations above indicate that thigh angular velocity must
be regulated to maintain a constant forward running speed. It has been
suggested that the limbs function as harmonic oscillators (McGeer, 1990),
which implies that ωc is related to the average of the absolute value of thigh
angular velocity during ωavg. Because of the continuous harmonic motion of
the thighs, it would be expected that a faster ωc would correspond to a
proportionately faster ωavg. From a mathematical standpoint, Eqns 7 and 8
imply that ωc and ωavg can be improved by increasing either f or A, or
increasing some combination of both (see examples in Fig. 8). Perhaps more
Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546
importantly, the equations above dictate that ωc and ωavg must both increase
for running speed to increase ( per hypothesis 1).
Enhanced ωavg should be related to an increased lower limb vertical
velocity at touchdown. As the front swinging thigh extends and causes limb
retraction during the last portion of the flight phase directly prior to impact
(e.g. Fig. 7A,B,D,E), faster angular velocity of the thigh should result in
faster tangential and vertical velocity of the lower limb in the distal portion
of the leg. Therefore, as ωavg increases in proportion to running speed, this
should also be associated with increases in thigh angular retraction velocity
(ωretr) prior to touchdown, and both of these measures should be positively
related to vertical velocity of the lower limb at touchdown across the range of
running speeds ( per hypothesis 2).
Subjects and participation
All testing and data collection were completed in one 90 min testing session at
the West Chester University of Pennsylvania laboratory and indoor athletic
facility. A total of 40 subjects volunteered and provided written informed
consent in accordance with the West Chester University of Pennsylvania
Institutional Review Board. This included 20 males (mean±s.e.m., age: 21.6±
0.5 years, height: 1.80±0.01 m, leg length: 0.94±0.01 m, mass: 79.7±3.0 kg)
and 20 females (age: 21.7±0.4 years, height: 1.67±0.02 m, leg length: 0.89±
0.01 m, mass: 59.0±1.4 kg). Male subjects included nine intercollegiate or
post-collegiate track and field athletes [≤400 m sprints (n=7), horizontal
jumps (n=2)], five intercollegiate team sport athletes [soccer (n=2), rugby
(n=2), American football (n=1)] and six recreationally trained subjects.
Female subjects included six intercollegiate or post-collegiate track and field
athletes [≤400 m sprints (n=5), 100 m hurdles (n=1)], eight intercollegiate
team sport athletes [gymnastics (n=3), soccer (n=2), rugby (n=1), softball
(n=1), basketball (n=1)], and six recreationally trained subjects. Complete
subject descriptive characteristics are listed in Table S1. Per inclusionary
criteria, all subjects were healthy and regularly active (defined as exercising
three or more times per week) at the time of testing.
Testing procedures
After subjects reviewed and signed consent forms, experimental procedures
were as follows. Subjects were provided with standardized compression
clothes and running track flats (Nike Waffle Racer version nine, Beaverton,
OR, USA). Subjects were measured for height using a standard measuring
tape and weighed on a digital scale (Supac Model EB-8008, Shanghai,
China). Subjects then performed a full-body warm up including jogging,
skipping, dynamic stretches and sub-maximal sprints. Next, for the purposes
of motion capture, subjects wore reflective markers placed on the heel and
ball of the foot on the lateral aspect of the running shoe, as well as on the
lateral aspect of the ankle, knee, hip and shoulder (lateral malleolus,
lateral femoral condyle, greater trochanter and acromial process, respectively).
There were 12 reflective markers total, six on each side of the body. A still
frame motion capture recording of each subject standing in the center of the
field of view was completed prior to the testing to serve as a reference for
kinematic analyses. Measurements of leg length from greater trochanter to
ground were determined from the still frame motion capture recording, with an
average leg-length value determined using measurements of right and left leg.
For the experimental testing, subjects completed 40 m running trials, and
data were captured and analyzed on four trials over a range of speeds.
Although the subjects were not directly paced by the investigators while
running, they were instructed to complete the trials at progressively faster
speeds, with the last trial being maximal. All trials were completed in a
running lane in an indoor athletic facility with a multi-purpose floor. The
running lane was 60 m in total length, allowing the subjects 20 m to safely
decelerate and stop after completing each 40 m running trial. Subjects
began each trial in an upright, ‘two-point’ stance with the preferred leg
Biology Open
forward and started at their own initiative. For the trials completed at less
than maximal intensity, subjects were instructed to gradually accelerate to
a cone placed at 25 m, and then to run at a constant speed from 25 m
through 40 m. For the maximal effort trials (interchangeably termed ‘top
speed’ trials), subjects were instructed to perform an all-out acceleration
from the beginning of the sprint, and continue at full speed all the way
through the finish line at the 40 m mark. Subjects were allowed complete
recovery between trials.
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Data collection and analysis
Three-dimensional kinematic data were recorded using an eight-camera
motion capture system collecting at 200 Hz (OptiTrack Prime 13 cameras
with Motive software from NaturalPoint, Corvallis, OR, USA). The field of
view captured by the motion capture system was eight meters in length, from
31–39 m in the running lane. This field of view was selected to ensure that
data could be collected for one complete gait cycle for even the tallest, fastest
subjects with the longest stride lengths. Prior to data collection, the capture
volume was dynamically calibrated using a wand with reflective markers at
known spacings. After data collection, motion capture files were exported
and uploaded to Microsoft Excel, where the data were up-sampled to
1000 Hz using linear interpolation and post-filtered using a low-pass,
fourth-order, zero-phase-shift Butterworth filter with a cutoff frequency of
25 Hz (Winter, 1990). Positional data from the 12 markers were used to form
a seven-segment model, including foot, shank and thigh on both legs, and a
head-arms-trunk segment (Winter, 1990).
In addition to measuring running speed using motion capture data
(described below), running speed was verified with a radar gun (Stalker ATS
II; Applied Concepts Inc., Plano, TX, USA). The radar data were collected at
47 Hz and exported to Microsoft Excel for analysis. Speed versus time data
were fit to a mono-exponential equation using an iterative least-squares
regression routine (Chelly and Denis, 2001; Samozino et al., 2016). Speed
versus time data were integrated to determine distance versus time, and then
used to calculate average speed from 31–39 m. Average running speed from
31–39 m, as determined by the radar data, showed a high level of agreement
compared to the motion capture method (R 2=0.99 and mean absolute error
<0.15 m s−1, see Appendix A and Fig. S1).
With regard to collecting top speed data in the field of view from 31–
39 m, prior research has illustrated that team-sport athletes approach
peak speed by 30 m into a sprint (Clark et al., 2019; Mendiguchia et al.,
2016). Although elite sprinters may not attain peak speed until 60 or
70 m into a 100 m dash (Krzysztof and Mero, 2013), split-time data from
100 m competitions has indicated that even the world’s fastest sprinters
have reached greater than 94% top speed by 30–40 m (Krzysztof and
Mero, 2013). Therefore, we deemed the 31–39 m field of view
appropriate to capture top speed for the heterogenous group of
subjects participating in this study. To assess the possibility that faster
subjects in this study were still accelerating through the motion capture
field of view during top speed trials, speed versus distance data from the
radar were examined to ensure that subjects ran at constant speed in the
motion capture zone. Constant speed was operationally defined as
changes in speed ≤0.3 m s−1 from 31–39 m, and all trials satisfied this
criterion.
Measurements
Based on the kinematic data, spatial–temporal and thigh angular kinematic
variables were quantified for all trials. Spatial-temporal variables included:
running speed, Tc, Tf, SR, SL, Lc and Ankle Vztd. All reported values are
trial averages from both right and left limbs during one complete gait cycle.
Instantaneous calculations of COM position and velocity were quantified
using the positions of the 12 reflective markers and segment information
from Winter (1990). Running speed was quantified from average COM
horizontal velocity in the 31–39 m field of view. With regards to
determining contact and flight time (Tc and Tf ), all trials were reviewed
using the motion capture software (NaturalPoint, Corvallis, OR, USA), with
instances of touchdown and takeoff determined from visual inspection of the
heel and ball marker vertical position relative to the running surface. This
method was validated compared to an in-ground laboratory force plate using
a separate sub-sample of subjects, and demonstrated a high level of
agreement with the force plate measurements (R 2=0.99 and Tc mean
absolute error <0.004 s, see Appendix A and Fig. S2). SR was calculated as
the inverse of the sum of ground contact time and flight time, or SR=1/
(Tc+Tf ). SL was calculated from running speed and step rate, or SL=Speed/
SR. Per Eqn 2 and Weyand et al. (2010), Lc was calculated by multiplying
running speed and ground contact time, or Lc=Speed×Tc. The absolute
value of vertical velocity of the ankle marker at the instant of touchdown
(Ankle Vztd) was used as a proxy for lower limb impact velocity ( per Clark
et al., 2017).
Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546
Additionally, several thigh angular kinematic variables were determined,
including: thigh angular position at touchdown (θtd), thigh angular position
at takeoff (θto), total thigh excursion during the ground contact phase (θc, the
sum of θtd and θto), peak thigh extension during flight (θext), peak thigh
flexion during flight (θflex), total thigh excursion from peak extension
through peak flexion (θtotal, the sum of θext and θflex), average thigh angular
retraction velocity measured from peak thigh flexion until touchdown ωretr,
average thigh angular velocity during the ground contact phase ωc, and
average thigh angular velocity during the entire gait cycle ωavg. Thigh
angular kinematics were quantified in an absolute frame of reference, with
the thigh angle compared to vertical, and not relative to trunk. Thigh angular
velocities (ωretr, ωc, and ωavg) were determined by calculating the derivative
of the thigh segment angular position versus time data, with angular
velocities measured as absolute values. As with the spatial–temporal
variables, reported thigh angular kinematic values were trial averages from
one complete gait cycle, with trial averages determined from both right and
left limbs.
Statistical analysis
To examine the relationship between running speed and the kinematic
variables of interest, the data were analyzed across all subjects and trials. Six
of the 40 subjects had sub-maximal trials with motion capture data that were
not usable due to marker occlusions. These trials were discarded, resulting in
n=154 total trials included in the data analysis. Across all subjects and trials,
the relationship between running speed and each of the kinematic variables
was evaluated using either simple linear regression or nonlinear regression
( power or second-order polynomial equations) to generate a best-fit
equation and coefficient of determination (R2), with x representing
running speed unless otherwise noted. Furthermore, differences in each of
the kinematic variables were analyzed across speeds with trials divided into
three categories based on percentage top speed, calculated for each subject
as a percentage of his or her fastest trial. These categories were slow (<75%
top speed, n=44 trials), intermediate (75 to 93%, n=48 trials), and fast
(>93%, n=62 trials). The normality of data was determined using the
D’Augustino and Pearson omnibus normality test or the Shapiro–Wilk test.
Parametric data were analyzed using 3x1 analysis of variance (ANOVA) and
Tukey’s post-hoc tests, and non-parametric data were analyzed using the
Kruskal–Wallis test (H statistic) and Dunn’s multiple comparisons tests.
In addition to the aforementioned analysis, top speed trials (n=40, one
trial per subject) were independently analyzed as a separate sub-set of data,
with simple linear regression used to determine the relationship between
each kinematic variable and top speed. To provide further insight into these
top speed trials, runners were grouped by sex (male and female) and top
speed (the faster ten subjects versus the slower ten subjects for each sex). For
each kinematic variable, separate independent t-tests were used to analyze
fast versus slow males and fast versus slow females. This analysis was
selected instead of a 2×2 ANOVA (sex by speed) because the primary
comparisons of interest were kinematic differences between fast and slow
runners analyzed within sex. For each data set, the normality of data was
determined using the D’Augustino and Pearson omnibus normality test
or the Shapiro–Wilk test. Parametric data were analyzed using unpaired
two-tailed t-tests, and non-parametric data were analyzed using
Mann–Whitney tests. For the top speed trials, absolute percentage
difference was also used to express the magnitude of difference between
fast versus slow group means for each variable, calculated as:
jmeanone  meantwo j
absolute percentage difference ¼  100:
fðmeanone þ meantwo Þ=2g
All data are expressed as mean±standard error of the mean (s.e.m.). The a
priori threshold for all significance tests was set at α=0.05. Statistical
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analyses of thigh angular kinematics were calculated in units of radians,
although for enhanced clarity, units of degrees are also presented where
relevant in Figs 2–7 and in Tables 1, 2, S2, S3. Power analyses for regression
and ANOVA were completed using G*Power (version 3.1.9, Kiel,
Germany), based on α=0.05, β=0.8 and moderate effect size. All other
statistics were completed using Microsoft Excel and GraphPad Prism
software (version 8, San Diego, CA, USA).
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Appendix A
Validation of running speed
Measurement of running speed using the motion capture method was
verified with a radar gun (Stalker ATS II; Applied Concepts Inc., Plano,
TX, USA). During all trials (n=154), the radar gun was placed on a tripod
at a height of 1.0 m, at a distance behind the starting line of 5.0 m. Radar
data were collected at 47 Hz, and then exported to Microsoft Excel for
analysis. Speed versus time data were fit with a mono-exponential equation
using an iterative least-squares regression routine (similar to Chelly and
Denis, 2001; Samozino et al., 2016). This equation models a runner’s
speed versus time curve as follows:
speedðtÞ ¼ speedmax ð1  eðt=tÞ Þ, ðA1Þ
where speedmax is the runner’s maximal speed and τ is the time constant.
Speed versus time data was then integrated to calculate distance versus
time:
ð Additional thigh angular kinematic results from all subjects and trials
distanceðtÞ ¼ speedðtÞdt, ðA2Þ
where distance is the horizontal position of the runner from the starting
line as a function of time. After distance versus time was determined,
average speed from 31–39 m was calculated. Average speed from 31–
39 m, as determined by radar data, showed a high level of agreement
compared to the motion capture method, with an absolute error of 0.11
±0.01 m s−1 (mean±s.e.m.). Average speed measured by the radar gun is
plotted against the motion capture method in Fig. S1.
Validation of ground contact time
To complete data analysis, it was necessary to develop an accurate method
for determining the instances of touchdown, takeoff, and total ground
contact time (Tc) using motion capture data. To accomplish this, a sub-
sample of subjects completed running trials over an in-ground laboratory
force plate with synchronized force and motion capture data. This sub-
sample consisted of ten subjects (mean±s.e.m.: seven males, age: 24.9±
2.3 years, height: 1.77±0.03 m, mass: 89.5±4.9 kg; three females, age:
21.0±0.6 years, height: 1.64±0.01 m, mass: 55.2±3.6 kg) who volunteered
and provided written informed consent in accordance with the local
university Institutional Review Board.
During a single 60 min testing session, subjects completed 12 trials in a
35 m laboratory runway with the force plate embedded at the 20 m point
of the runway. All procedures related to subject clothing/footwear,
measurements, warm-up, and motion capture marker placement were
identical to those described in the Materials and Methods. Although subjects
were not directly paced by the investigators while running, they were
instructed to complete the trials at progressively faster speeds, with the last
three trials being maximal. Subjects lined up in an upright stance 20 m
behind the force plate and, at their own initiative, ran forward and stepped on
the force plate during the course of a running stride. Subjects were instructed
to run naturally as they ran over the force plate. If the subject missed or
partially struck the force plate, or visibly altered their mechanics to strike the
plate, the data obtained from the trial were discarded. A total of 120 total
trials were completed, yielding 95 valid trials with synchronized force and
motion capture data where the subject fully struck the force plate without
altering normal running mechanics.
Data were synchronously collected with an in-ground force plate (Kistler
5691A, 0.6 m×0.9 m, Winterthur, Switzerland) and an eight-camera motion
capture system (OptiTrack Prime 13, Corvallis, OR, USA). Cameras were
placed around the force plate and provided a 6 m field of view centered on the
force plate. Prior to data collection, the capture volume was dynamically
calibrated using a wand with reflective markers at known spacings. The force
plate data were collected at 1000 Hz and the motion capture data at 200 Hz.
Force plate data were post-filtered in Microsoft Excel using a low-pass, fourth-
order, zero-phase-shift Butterworth filter with a cutoff frequency of 25 Hz
(Winter, 1990). All motion capture files were analyzed in an identical manner
to that presented in the Materials and Methods. Additionally, trials were
recorded using a high-speed video camera (Apple iPad 9.7, Apple USA,
Cupertino, CA, USA) to ensure that the foot fully contacted the force plate and
that no part of the foot landed off the force plate.
Biology Open (2020) 9, bio053546. doi:10.1242/bio.053546
Kinematic data were analyzed to determine the spatial and temporal variables
associated with touchdown and takeoff. All trials were reviewed using the
motion capture software (NaturalPoint, Corvallis, OR, USA), with instances of
touchdown and takeoff (and thus total Tc) determined from visual inspection of
the heel and ball marker vertical position relative to the running surface. This
visual inspection method was validated compared to the force plate, which was
used as the standard reference for measuring ground contact time (defined as the
time when vertical force measured by the force plate exceeded 20 N). Compared
to force plate data, this kinematic visual inspection method determined
touchdown with a mean absolute error of 2.1±0.2 ms, takeoff with a mean
absolute error of 2.7±0.2 ms, and Tc with a mean absolute error of 3.4±
0.3 ms (mean±s.e.m.). Contact times measured by the force plate are
plotted against those determined by the motion capture visual inspection
method in Fig. S2.
Appendix B
Increases in θtotal across the range of speeds were due to both increases in
θflex (θflex=0.102x+0.273, R 2=0.51, P<0.0001; Table S2: H=45.2,
P<0.0001) and increases in θext [θext=0.020x+0.367, R 2=0.06, P=0.003;
Table S2: F(2,151)=16.0, P<0.0001], though increases in θext from
intermediate to fast speeds were not significant (Table S2: P=0.18).
Increases in θc across the range of speeds were due to both increases in θtd
[θtd=0.033x+0.335, R 2=0.33, P<0.0001; Table S2: F(2,151)=29.5,
P<0.0001] and increases in θto (θto=0.30x 0.16, R 2=0.03; Table S2:
H=13.63, P<0.01), though increases in θto from intermediate to fast
speeds were not significant (Table S2: P>0.9).
Additional thigh angular kinematic results from top speed trials
Across all top speed trials, θflex was positively and significantly related to
faster top speeds (θflex=0.132x−0.023, R2=0.38, P<0.0001), although
differences in θflex did not reach significance when analyzing fast versus
slow runners within sex (Table S3, males: P=0.062, Δ=16.5%; females:
P=0.059, Δ=13.0%). Across all top speed trials, θext was negatively and
significantly related to faster top speeds (θext=−0.057x+1.044, R 2=0.21,
P=0.003), although differences in θext did not reach significance when
analyzing fast versus slow runners within sex (Table S3, males: P=0.117,
Δ=16.7%; females: P=0.51, Δ=5.1%). Across all top speed trials, θtd was not
significantly related to top speed (θtd=0.018x+0.459, R2=0.05, P=0.159) or
when analyzing fast versus slow runners within sex (Table S3, males: P>0.9,
Δ=0.0%; females: P>0.9, Δ=0.1%). Across all top speed trials, θto
was negatively and significantly related to faster top speeds
(θto=−0.043x+0.817, R 2=0.21, P=0.003), although differences in θto did
not reach significance when analyzing fast versus slow runners within sex
(Table S3, males: P=0.09, Δ=15.7%; females: P=0.37, Δ=7.4%).
Acknowledgements
The authors thank Tyler Whitacre, Sabrina Mangeri, Samantha Jenks, Sean
Leonard, Ronald Mungin, Andrew Slater, Nick Jachwak, Kyle McCormick, Dylan
Ferron, and Marshall Lane. The authors also thank the subjects for their
participation, especially the WCUTF program and Jason Kilgore.
Competing interests
The authors declare no competing or financial interests.
Author contributions
Conceptualization: K.P.C.; Methodology: K.P.C.; Validation: K.P.C., C.R.M.; Formal
analysis: K.P.C., C.R.M., D.J.S.; Investigation: K.P.C., C.R.M., D.J.S.; Data curation:
K.P.C., C.R.M., D.J.S.; Writing - original draft: K.P.C., C.R.M., D.J.S.; Writing - review
& editing: K.P.C., C.R.M., D.J.S.; Supervision: K.P.C., D.J.S.; Project administration:
K.P.C., D.J.S.
Biology Open
Funding
This research received no specific grant from any funding agency in the public,
commercial or not-for-profit sectors.
Supplementary information
Supplementary information available online at
https://bio.biologists.org/lookup/doi/10.1242/bio.053546.supplemental
12
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